Cerebral asymmetries on magnetic resonance imaging

Magnetic Resonance Imaging (MRI) reveals anatomical asymmetries of the sulcal demarcation of the posterior operculum from the parietal cortex that correlate with handedness in normal subjects. The sulcal demarcation is greater on the right in most right handers, but evenly distributed in left handers. This in vivo asymmetry corresponds to the commonly larger planum temporale on the left side. Hand Preference, Hand Performance, Dichotic Listening and Visual Field Preference measures were correlated with several measures of anatomical asymmetries seen on MRI. The sulcal demarcation, anterior frontal width and parietal width correlated best with hand performance, as measured on a dot tapping task. The biological significance of the distribution of anatomical and functional asymmetries is discussed.     

Muscular or directional preferences in finger movement as a function of handedness

Right handed, left handed, and "ambiguous" male and female subjects performed circular index finger movements. Analysis of preferred direction of movements showed that strong right handers tend to move both left and right index fingers in the same direction, and familial left handers tend to move them in opposite directions. Since right handers tend to possess a strongly dominant left hemisphere, while familial left handers exhibit a high degree of cerebral lateral equipotentiality, the result indicates that interhemispheric interference in a motor skill consists of activation of inappropriate muscles of the non-preferred hand by the dominant ipsilateral hemisphere as its attempts to force that hand to conform to the direction of movement preferred by the dominant hand.     

Hemispheric asymmetries of motor versus nonmotor processes during (visuo)motor control

Language and certain aspects of motor control are typically served by the left hemisphere, whereas visuospatial and attentional control are lateralized to the right. Here a (visuo)motor tracing task was used to identify hemispheric lateralization beyond the general, contralateral organization of the motor system. Functional magnetic resonance imaging (fMRI) was applied in 40 male right‐handers (19–30 yrs) during line tracing with dominant and nondominant hand, with and without visual guidance. Results revealed a network of areas activating more in the right than left hemisphere, irrespective of the effector. Inferior portions of frontal gyrus and parietal lobe overlapped largely with a previously described ventral attention network responding to unexpected or behaviourally relevant stimuli. This demonstrates a hitherto unreported functionality of this circuit that also seems to activate when spatial information is continuously exploited to adapt motor behaviour. Second, activation of left dorsal premotor and postcentral regions during tracing with the nondominant left hand was more pronounced than that in their right hemisphere homologues during tracing with the dominant right hand. These activation asymmetries of motor areas ipsilateral to the moving hand could not be explained by asymmetries in skill performance, the degree of handedness, or interhemispheric interactions. The latter was measured by a double‐pulse transcranial magnetic stimulation paradigm, whereby a conditioning stimulus was applied over one hemisphere and a test stimulus over the other. We propose that the left premotor areas contain action representations strongly related to movement implementation which are also accessed during movements performed with the left body side. Hum Brain Mapp, 2011. © 2010 Wiley‐Liss, Inc.     

Neurostructural correlates of consistent and weak handedness

ABSTRACT

Various cognitive differences have been reported between consistent and weak handers, but little is known about the neurobiological factors that may be associated with this distinction. The current study examined cortical structural lateralization and corpus callosum volume in a large, well-matched sample of young adults (N?=?164) to explore potential neurostructural bases for this hand group difference. The groups did not differ in corpus callosum volume. However, at the global hemispheric level, weak handers had reduced or absent asymmetries for grey and white matter volume, cortical surface area, thickness, and local gyrification, relative to consistent handers. Group differences were also observed for some regional hemispheric asymmetries, the most prominent of which was reduced or absent gyrification asymmetry for weak handers in a large region surrounding the central sulcus and extending into parietal association cortex. The findings imply that variations in handedness strength are associated with differences in structural lateralization, not only in somatomotor regions, but also in areas associated with high level cognitive control of action.     

Asymmetries in motor attention during a cued bimanual reaching task: Left and right handers compared

Several studies have indicated that right handers have attention biased toward their right hand during bimanual coordination ( [Buckingham and Carey, 2009] and [Peters, 1981] ). To determine if this behavioral asymmetry was linked to cerebral lateralization, we examined this bias in left and right handers by combining a discontinuous double-step reaching task with a Posner-style hand cueing paradigm. Left and right handed participants received a tactile cue (valid on 80% of trials) prior to a bimanual reach to target pairs. Right handers took longer to inhibit their right hand and made more right hand errors, suggesting that their dominant hand was more readily primed to move than their non-dominant hand, likely due to the aforementioned attentional bias. Left handers, however, showed neither of these asymmetries, suggesting that they lack an equivalent dominant hand attentional bias. The findings are discussed in relation to recent unimanual handedness tasks in right and left handers, and the lateralization of systems for speech, language and motor attention.     

Evidence of a possible isolation of left hemisphere visual and motor areas in sinistrals employing an inverted handwriting posture

Levy's hypothesis of ipsilateral motor control of the lateral distal musculature in left handers employing an inverted handwriting posture was tested in a true simple reaction time paradigm. Left-handers employing a non-inverted handwriting posture (NHP) showed performances consistent with normal (contralateral) motor control organization. Inverted handwriting posture (IHP) sinistrals showed performances that were not consistent with Levy's hypothesis nor with the simple hypothesis of normal (contralateral) control. The results for IHP sinistrals were compatible with a model of stimulus-response mediation which posits contralateral motor control and an intrahemispheric “disconnection” of left hemisphere visual areas from left hemisphere manual motor areas. The hypothesis that IHP sinistrals must utilize interhemispheric pathways for motor responses to stimuli channeled to, or formulated in, the left hemisphere was discussed in relation to the hypothesis of topographic inversion consequent to transcallosal relay. The proposed model suggests that hand inversion in writing is an accommodation to the consequences of interhemispheric relay and not to ipsilateral motor control.     

Mapping hemispheric asymmetries of the macaque cerebral cortex during early brain development

Studying cortical hemispheric asymmetries during the dynamic early postnatal stages in macaque monkeys (with close phylogenetic relationship to humans) would increase our limited understanding on the possible origins, developmental trajectories, and evolutional mechanisms of brain asymmetries in nonhuman primates, but remains a blind spot to the community. Via cortical surface‐based morphometry, we comprehensively analyze hemispheric structural asymmetries in 134 longitudinal MRI scans from birth to 20 months of age from 32 healthy macaque monkeys. We reveal that most clusters of hemispheric asymmetries of cortical properties, such as surface area, cortical thickness, sulcal depth, and vertex positions, expand in the first 4 months of life, and evolve only moderately thereafter. Prominent hemispheric asymmetries are found at the inferior frontal gyrus, precentral gyrus, posterior temporal cortex, superior temporal gyrus (STG), superior temporal sulcus (STS), and cingulate cortex. Specifically, the left planum temporale and left STG consistently have larger area and thicker cortices than those on the right hemisphere, while the right STS, right cingulate cortex, and right anterior insula are consistently deeper than the left ones, partially consistent with the findings in human infants and adults. Our results thus provide a valuable reference in studying early brain development and evolution.     

Assessment of sensorimotor cortical representation asymmetries and motor skills in violin players

As a model for use-dependent plasticity, the brains of professional musicians have been extensively studied to examine structural and functional adaptation to unique requirements of skilled performance. Here we provide a combination of data on motor performance and hand representation in the primary motor and somatosensory cortex of professional violin players, with the aim of assessing possible behavioural consequences of sensorimotor cortical asymmetries. We studied 15 healthy right-handed professional violin players and 35 healthy nonmusician controls. Motor and somatosensory cortex asymmetry was assessed by recording the motor output map after transcranial magnetic stimulation from a small hand muscle, and by dipole source localization of somatosensory evoked potentials after electrical stimulation of the median and ulnar nerves. Motor performance was examined using a series of standardized motor tasks covering different aspects of hand function. Violin players showed a significant right-larger-than-left asymmetry of the motor and somatosensory cortex, whereas nonmusician controls showed no significant interhemispheric difference. The amount of asymmetry in the motor and somatosensory cortices of musicians was significantly correlated. At the behavioural level, motor performance did not significantly differ between musicians and nonmusicians. The results support a use-dependent enlargement of the left hand representation in the sensorimotor cortex of violin players. However, these cortical asymmetries were not paralleled by accompanying altered asymmetries at a behavioural level, suggesting that the reorganisation might be task-specific and does not lead to improved motor abilities in general.     

Network asymmetry of motor areas revealed by resting-state functional magnetic resonance imaging

There are ample functional magnetic resonance imaging (fMRI) studies on functional brain asymmetries, and the asymmetry of cerebral network in the resting state may be crucial to brain function organization. In this paper, a unified schema of voxel-wise functional connectivity and asymmetry analysis was presented and the network asymmetry of motor areas was studied. Twelve healthy male subjects with mean age 29.8 ± 6.4 were studied. Functional network in the resting state was described by using functional connectivity magnetic resonance imaging (fcMRI) analysis. Motor areas were selected as regions of interest (ROIs). Network asymmetry, including intra- and inter-network asymmetries, was formulated and analyzed. The intra-network asymmetry was defined as the difference between the left and right part of a particular functional network. The inter-network asymmetry was defined as the difference between the networks for a specific ROI in the left hemisphere and its homotopic ROI in the right hemisphere. Primary motor area (M1), primary sensory area (S1) and premotor area (PMA) exhibited higher functional correlation with the right parietal-temporal-occipital circuit and the middle frontal gyrus than they did with the left hemisphere. Right S1 and right PMA exhibited higher functional correlation with the ipsilateral precentral and supramarginal areas. There exist the large-scale hierarchical network asymmetries of the motor areas in the resting state. These asymmetries imply the right hemisphere dominance for predictive motor coding based on spatial attention and higher sensory processing load for the motor performance of non-dominant hemisphere.     

Effects of stimulation intensity, gender and handedness upon auditory evoked potentials.

Left handers and women show less anatomical brain asymmetry, larger corpus callosum and more bilateral representation of specific functions. Sensory and cognitive components of cortical auditory evoked potentials (AEP) have been shown to be asymmetric in right handed males and to be influenced by stimulus intensity. In this study the influence of sex, handedness and stimulus intensity upon AEP components is investigated under basal conditions of passive attention. 14 right handed males, 14 right handed females, 14 left handed males, and 14 left handed females were studied while lying awake and paying passive attention to auditory stimulation (series of 100 binaural clicks, duration 1 msec, rate 1/sec, at four intensities). Cz, C3 and C4 referenced to linked mastoids and right EOG were recorded. Analysis time was 400 msec, average evoked potentials were based on 100 clicks. Stimulus intensity and gender affect early sensory components (P1N1 and N1P2) at central leads, asymmetry is influenced only by handedness, right handers showing larger P1N1 amplitudes over the right hemisphere.     

A functional magnetic resonance imaging study of cortical asymmetry in bipolar disorder

Objectives:  Individuals with bipolar disorder (BPD) exhibit motor, perceptual, and cognitive disturbances involving predominantly right hemisphere dysfunction. This asymmetry has been used to advance the hypothesis that the pathogenesis of bipolar disorder may be related to disturbances of the right cerebral hemisphere. We employed functional magnetic resonance imaging to examine hemispheric asymmetries in manic and depressed BPD. A secondary goal of the study was to examine effects of psychotropic medications on blood volume changes in the motor cortices.
Methods:  We studied 18 right-handed BPD and 13 right-handed normal healthy comparison subjects. Blood oxygen level dependent (BOLD) responses in the primary motor area (M1) and supplementary motor area (SMA) of both hemispheres were elicited during reaction time (RT) tasks.
Results:  Healthy subjects activated the SMA in a reciprocal fashion with significantly greater activity in the left SMA for right hand trials and the right SMA for left hand trials. Depressed BPD subjects failed to show this normal reciprocity indicating a failure to suppress unwanted activity in the ipsilateral right SMA, whereas manic BPD subjects failed to suppress unwanted ipsilateral SMA activity in both hemispheres. Manic and depressed BPD subjects exhibited greater activity in the left primary motor area suggesting increased cortical excitability. BPD subjects treated with antipsychotics or mood-stabilizing medications exhibited longer RTs, lower BOLD responses in M1 and SMA, and a loss of normal hemispheric asymmetry in the SMA than untreated subjects.
Conclusions:  The presence of a right hemisphere disturbance in BPD is consistent with the hypothesis that the right hemisphere may be dominant in mood regulation. The presence of both left and right hemisphere disturbances in mania may explain the coexisting psychotic and affective symptoms observed in this condition.     

Asymmetries in the cerebral dimensions and fissures of the dog

The surface dimensions and fissures of the right and left cerebral hemispheres were compared in dogs. The right hemisphere was significantly larger in length and height than the left hemisphere; there was no significant difference in the width of the right and left hemispheres. The mean length of the cruciate sulcus did not significantly differ on the right and left sides except individual for asymmetries. The difference in the mean length of the right and left Sylvian fissures was not significant, but the right Sylvian fissure was significantly lower than the left. The base of the right planum temporale tended to be larger than the left. These anatomical asymmetries did not correlate with paw preference. The data are consistent with notion that asymmetrical patterns cannot be a distinctive feature of the human brain.     

Is interhemispheric transfer of visuomotor information asymmetric? Evidence from a meta-analysis

Using a meta-analytic procedure we have analysed 16 studies employing a simple unimanual reaction time (RT) paradigm and lateralized visual stimuli to provide an estimate of interhemispheric transfer time in normal right-handed subjects. We found a significant overall RT advantage of the left visual field over the right and of the right hand over the left. These asymmetries can be explained by a superiority of the right hemisphere for the detection of simple visual stimuli and by a corresponding superiority of the left hemisphere for the execution of the manual response, respectively. Alternatively, they may be interpreted as related to an asymmetry of interhemispheric transmission of visuomotor information, with transfer from the right hemisphere (side of stimulus entry) to the left (side of response generation) faster than in the reverse direction. Although a direct test of these hypotheses is still lacking, we think that the evidence available is more in keeping with the latter possibility.     

Sex differences in line bisection as a function of hand.

If subjects are asked to indicate the midpoint of a horizontal line, they tend to bisect it left of the center, a phenomenon called 'pseudoneglect'. Assuming that this task evokes visuospatial processes, the left bias is generally considered to arise from a right-hemispheric activation. Numerous factors affect pseudoneglect. Although, only few studies have examined the influence of sex in visual line bisection, most of theses studies reported no significant main effect of sex. Possible interactions between sex and other performance factors that are linked to the motor component of this task, e.g. hand use, are rarely examined. We studied the interaction of these two variables using right-handed females and males in a line bisection task. The results clearly indicate that hand use and sex interact, with females showing the left bias to a similar extent with both hands while males show the bias predominantly with the left hand. Moreover, the position of the lines (left, middle and right) significantly affect left bias in visual line bisection and interact with hand use. It is hypothesized that the larger cross-section of the posterior corpus callosum in females enables a stronger interhemispheric connectivity of visuospatial cortical areas resulting in a strong left-sided bias in hand motor cortical areas of both hemispheres. In males, motor cortical activation would accordingly be mainly restricted to the right hemisphere.     

Interhemispheric differences of hand muscle representation in human motor cortex

Focal magnetic transcranial stimulation (TCS) is employed for mapping of the motor cortical output to abductor digiti minimi (ADM) muscle. The aim of this study was to evaluate the interhemispheric asymmetries in normals. Motor maps were obtained through motor evoked potentials (MEPs) recordings from ADM muscle in 20 healthy subjects in right and left hemispheres TCS. Measurement of several indexes such as excitability threshold, MEPs amplitude, MEPs latency, and silent period duration did not show differences between the hemispheres. Moreover, no interhemispheric asymmetries were found when the amplitude ratio values were analyzed. The hand motor cortical area, as represented by the number of responsive sites (3.6 vs. 3.5) and the “hot spot” site localization presented a fairly symmetrical organization. Absolute values displayed a relatively wide intersubject variability, while their interhemispheric differences were extremely restricted. This observation can offer a new tool in diagnosing and following up neurological disorders affecting the central motor system, mainly for those concerning monohemispheric lesions. © 1997 John Wiley & Sons, Inc. Muscle Nerve, 20, 535–542, 1997.     

Preferred hand and familial sinistrality in dichotic monitoring

Ear asymmetries in dichotic monitoring performance were assessed in 105 subjects (77M, 28F). A right ear advantage (REA) was found more often in right than mixed or left handed subjects. Handedness did not affect the magnitude of asymmetry. Familial sinistrality could only be examined in male left handers, and was found to increase the incidence of REA's. Non-familial sinistrals were equally distributed in terms of left, right and no ear advantage.     

Hemispheric asymmetry for trajectory perception

Broadly, the right hemisphere is known to be specialized for spatial processing whereas the left hemisphere is known to be specialized for temporal processing. However, it remains unclear how both hemispheres interact when processing spatio-temporal information. This study investigates, from a behavioral point of view, whether spatio-temporal processing involved in trajectory perception generates hemispheric asymmetries. An experimental task requiring the prediction of coincidence between ballistic trajectories and a stationary target was used. Reaction times were analyzed according to various interhemispheric conditions determined by the visual hemifield on which the stimulus was presented and the hand of response. There was shorter reaction time for the left hand than the right hand, and shorter reaction times for the left visual hemifield than the right visual hemifield for both hands. From these findings, it is inferred that there is likely to be right hemisphere specialization for trajectory perception and that this hemispheric asymmetry is independent of handedness.     

Semantic processing in the left versus right cerebral hemispheres following unilateral hand contractions

Unilateral hand contractions increase activation in the motor cortex of the contralateral hemisphere, providing a means to alter the relative degree of activation in the right hemisphere versus the left hemisphere through spreading activation. Prior research reported enhanced verbal creativity as measured by performance on remote associate problems in Hebrew from left-hand contractions (right-hemisphere activation). We sought to extend the previous findings to English problems and to homograph interpretation. In Experiment 1, unilateral hand contractions in fact altered performance on the English remote associates, but in the direction of improved performance following right-hand contractions and left-hemisphere activation. In Experiment 2, the probability of retrieving atypical interpretations of homographs with multiple meanings was least likely for left-hemisphere dominant strong right handers, but the hand contraction manipulation had no effect.     

Interrelations between motivational stance, cortical excitability, and the frontal electroencephalogram asymmetry of emotion: a transcranial magnetic stimulation study

Several electrophysiological studies have provided evidence for the frontal asymmetry of emotion. In this model the motivation to approach is lateralized to the left, whereas the motivation to avoidance is lateralized to the right hemisphere. The aim of the present experiment was to seek evidence for this model by relating electrophysiological and phenomenological indices of frontal asymmetry to a direct measure of cortical excitability. Frontal asymmetrical resting states of the electroencephalogram and motivational tendencies indexed by the Carver and White questionnaire were compared with neural excitability of the left and right primary motor cortex as assessed by transcranial magnetic stimulation in 24 young healthy right-handed volunteers. In agreement with the model of frontal asymmetry, predominant left over right frontal cortical excitability was associated with enhanced emotional approach relative to emotional avoidance. Moreover, the asymmetries of brain excitability and approach-avoidance motivational predispositions were both reflected by frontal beta (13-30 Hz) electroencephalogram asymmetries. In conclusion, the currently demonstrated interconnections between cortical excitability, electrophysiological activity, and self-reported emotional tendencies for approach or avoidance support the frontal asymmetry of emotion model and provide novel insights into its biological underpinnings.     

Histological asymmetries of primary motor cortex predict handedness in chimpanzees (Pan troglodytes)

Like humans, chimpanzees display robust and consistent hand preferences during the performance of certain tasks. Although correlations have been demonstrated between gross anatomic measures of primary motor cortex asymmetry and handedness in captive chimpanzees, the relationship between histological architecture and behavioral lateralization has not yet been investigated. Therefore, we examined interhemispheric asymmetry of several different microstructural characteristics of the primary motor cortex in the region of hand representation from 18 chimpanzees tested on a coordinated bimanual task before death. At the population level our data showed leftward bias for higher layer II/III neuron density. Of note, however, there was no population-level asymmetry in the areal fraction of Nissl-stained cell bodies, a finding that differs from previous studies of this cortical region in humans. Nonetheless, we found that asymmetry in the density of layer II/III parvalbumin-immunoreactive interneurons was the best predictor of individual hand preference. These results suggest that histological asymmetries are related to handedness in chimpanzees, while overall patterns of asymmetry at the population level might differ from humans.